Accepted Manuscript

Amino acid composition of flesh-coloured potatoes as affected by storage con-

ditions

Anna Pęksa, Joanna Miedzianka, Agnieszka Nemś

PII: S0308-8146(18)30998-1
DOI: https://doi.org/10.1016/j.foodchem.2018.06.026
Reference: FOCH 22987

To appear in: Food Chemistry

Received Date: 8 January 2018

Revised Date: 25 May 2018
Accepted Date: 5 June 2018

Please cite this article as: Pęksa, A., Miedzianka, J., Nemś, A., Amino acid composition of flesh-coloured potatoes
as affected by storage conditions, Food Chemistry (2018), doi: https://doi.org/10.1016/j.foodchem.2018.06.026

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 Amino acid composition of flesh-coloured potatoes as affected by storage conditions

Anna Pęksa, Joanna Miedzianka,* Agnieszka Nemś

Department of Food Storage and Technology, Faculty of Biotechnology and Food Science,

Wroclaw University of Environmental and Life Sciences

*Corresponding author: Joanna Miedzianka, Department of Food Storage and Technology,

Faculty of Biotechnology and Food Science, Wroclaw University of Environmental and Life

Sciences, Poland

Email: joanna.miedzianka@upwr.edu.pl
Abstract

The study determined nitrogen compounds and amino acid profile in dry matter of potatoes

differing in flesh colour, stored at 2 °C and 5 °C for three and six months. With increased storage

time, the total protein content and particularly amino acid content declined. The coagulable protein

content increased at three months’ storage by 25%. The majority of the amino acid content

decreased from 19 to 6% and from 38 to 21% after three and six months’ storage, respectively.

Storage temperature did not influence the coagulable protein content or serine, glycine, cysteine,

tyrosine and phenylalanine. However, potatoes stored at 2 ºC contained slightly more amino acids

than tubers stored at 5 ºC. Independently of the storage conditions, potatoes of yellow-fleshed

Fresco and red-fleshed Herbie 26 varieties were characterised by a relatively high nutritive value,

limited by leucine (CS = 84), methionine plus cysteine (CS = 78) and leucine (CS = 72),

respectively.

Keywords: coloured potatoes, storage, amino acids profile, protein quality

1. Introduction

Potatoes are very popular as an inexpensive food product, available throughout the year due to

their suitability for long-term storage. These vegetables are a valued raw material in starch and

alcohol manufacture, for instance, and a valuable consumer product, mainly for their versatility of

usage and favourable sensory and nutritional properties. Moreover, potatoes outperform other

products, such as wheat, rice or corn, in terms of nutritional value, cost of cultivation and storage

(Friedman, 1996; Lister & Munro, 2000).

An increasing number of studies have described potato varieties with coloured flesh, notably

purple and red. Despite similarities in chemical composition to traditional, yellow, creamy or white

varieties (Jansen & Flamme, 2006; Lachman et al., 2012; Pęksa et al., 2013), coloured-fleshed

potatoes contain anthocyanins, polyphenolic compounds with beneficial effects on human health.

Tubers with purple or red flesh can be a profitable source of anthocyanins, similar to cranberries

and superior to red cabbage. These compounds, besides displaying antioxidant properties, exhibit

the activity in reducing the risk of chronic diseases of the nervous system; furthermore, they give

tubers interesting functions not found in potatoes of traditional light colour. Potatoes with coloured

flesh are similar to traditional fleshed tubers in terms of the content of nitrogen compounds (Jansen

& Flamme, 2006). However, as has been shown in previous work (Pęksa et al., 2013), leucine is the

amino acid limiting the quality of protein contained in purple- and red-fleshed varieties, whereas, in

yellow-fleshed cultivars, it is primarily the sulfur amino acids, methionine and cysteine.

Coloured-flesh potatoes cultivars allow long-term storage without notable loss of anthocyanins

and belong to the so-called low-cost crops; their production and storage are well established.

Coloured-flesh potatoes exhibit lower levels of anti-nutritional compounds, like glycoalkaloids,

compared to yellow- and cream-fleshed varieties (Tajner-Czopek, Rytel, Kita, Pęksa, & Hamouz,

2012).

Regardless of flesh colour, potatoes contain protein of high nutritive value which consists of

two main fractions: coagulable protein (nitrogen compounds precipitable with trichloroacetic acid)
and non-protein organic compounds, such as free amino acids. This coagulable protein is a valuable

foodstuff because of its well-balanced amino acid composition (van Gelder & Vonk, 1980). Potato

protein is of great biological and nutritional value, comparable with egg white, and its chemical

score (CS) ranges from 57 to 69 (Kapoor, Desbrough, & Li, 1975; Mitrus, Stankiewicz, Steć,

Kamecki, & Starczewski, 2003; Pęksa, 2003; Pęksa, Rytel, Kita, Lisińska, & Tajner-Czopek, 2009).

Potatoes contain significant amounts of aspartic and glutamic acids and their amides, as well as

essential amino acids (EAAs) like leucine, lysine, phenylalanine, valine and tyrosine (Burton, 1989;

Zimnoch-Guzowska & Flis, 2006; Pranaitiene, Danilcenko, Jariene, & Dabkevicius, 2008).

About 50% of the total nitrogen presented in potatoes is derived from proteins (Eppendorfer,

Eggum, & Bille, 1979; Kapoor et al., 1975); around 40% of the remaining soluble non-protein

nitrogen consists of the above-mentioned free amino acids and their amides, while 10% comprises

non-protein nitrogen associated with glycoalkaloids, some vitamins, purines, pyrimidines and

secondary metabolites (Friedman, 1996). About 35% of the soluble protein is glycoprotein of 44–45

kDa molecular weight, known as patatin or tuberin, 25% of the protein includes protease inhibitors,

and the remaining 40% contains other proteins with different properties (Deveaux-Gobert, 2008;

Pęksa et al., 2009).

Apart from their nutritional functions, amino compounds, such as amino acids, peptides and

proteins, exhibit antioxidant activity and thus are considered important in plants and animals.

Antioxidant amino acids include methionine, cysteine, tryptophan, tyrosine, histidine, lysine and

proline.

Storage of potatoes is aimed at extending the shelf-life while minimising quantitative and

qualitative losses. The suitability of potatoes for long-term storage is associated with the genetic

properties of the variety, which can be changed under the influence of the growing conditions and

storage. Storage stability of varieties depends on the resting period of the tubers, the intensity of life

processes occurring in the tubers, and the resistance to mechanical damage and susceptibility of

tubers to fungal and bacterial diseases during vegetation and storage (Kołodziejczyk, 2016). During
storage, the chemical composition of potatoes is changed, mainly by the temperature. At increased

storage temperature, respiration, transpiration and germ growth intensify, causing apparent

increases in the dry matter (DM) content and loss of reducing sugars and starch. Most potato

varieties exhibit low life activity when stored at 4–6 ºC. Table potatoes are usually stored at about

4ºC, which extends the period of dormancy, reduces the intensity of the growth of germs, stabilises

the DM content of tubers and lowers the natural losses, also limiting the development of the

majority of storage diseases (Czerko, Zgórska, & Grudzińska, 2012). However, this leads to the

accumulation of reducing sugars, along with protein degradation, which is consistent with an

increase in proteolytic enzyme activity, enhanced by low-temperature conditions (Brierley, Bonner,

& Cobb, 1996). Both of these processes may act as determinants of potato processing quality. The

research of various authors shows that long-term storage results in a decrease of the content of most

amino acids and a similar downward trend can be seen in total protein (Brierley et al., 1996; Černá

& Kráčmar, 2010). In these studies, it is also pointed out that amino acid composition of potatoes

during long-term storage, usually, in low temperature conditions, depends primarily on the time of

storage but also on the potato variety.

The content and structure of nitrogen compounds is modified in yellow-fleshed varieties during

long-term storage of potato tubers (Galdón, Mesa, Rodriquez, & Romero, 2010; Jansen & Flamme,

2006; Rexen, 1976), while potatoes with a coloured flesh in this regard are proportionally less

tested. There is no information in published literature about the impact of the presence of

anthocyanins in potato tubers on the variations in the content and composition of nitrogen

compounds in potatoes during storage. It is worth learning the factors influencing the

transformation of nitrogen compounds in potato varieties with red and purple flesh as affecting their

nutritional value, both due to the growing interest of consumers and potato producers, but also due

to the extensive research on varieties with coloured flesh in terms of their traits as raw materials in

the food industry and dietetic food. The aim of this study was to investigate the magnitude and

direction of changes in the content of nitrogen compounds in total, protein nitrogen, and amino
acids, and thus the nutritional value of potatoes differing in varietal characteristics, including the

colour of flesh, originating from different growing conditions, during long-term storage at low

temperature.

2. Materials and methods

2.1. Raw material

Six varieties of potato cultivated in the year 2014 were studied, including purple-fleshed

(Herbie 26 and Rote Emma), red-fleshed (Blaue Annelise and Blue Congo) and traditional yellow-

fleshed (Vineta and Fresco). The coloured-fleshed potatoes were grown in the test field at the

station of The Central Institute for Supervising and Testing in Agriculture at Přerov nad Labem (the

Czech Republic) and potatoes of the traditional yellow-fleshed varieties of Vineta and Fresco were

obtained from a potato producing plant in Lower Silesia in Poland. The samples of potato tubers

were harvested after reaching full maturity. Average laboratory samples of 20 kg tubers of each

colour-fleshed variety were selected randomly from the collected field samples (40-50 kg).

Mechanically damaged and green potatoes were rejected. Thereafter, the 20 kg samples of tubers of

each variety were divided into two repetitions. Each sample of 10–15 tubers (weighing

approximately 1.5 kg) was stored concurrently in paper bags for zero (at start of storage), three and

six months at two low temperatures (2 ºC and 5 ºC), exposed to the air, at constant relative humidity

(85% ±2%; thermohydrometer TH-130; Hama, Mannheim, Germany). After each storage period,

the potatoes were analysed. Prepared material was stabilised by lyophilisation and stored below ‒18

ºC in closed polyethylene tubs for further analysis.

2.2. Proximate analysis

The DM, starch, and total and coagulable nitrogen content were evaluated according to the

Association of Analytical Chemists’ method (AOAC, 1995). Protein content was calculated using a

conversion factor of 6.25.

2.3. Determination of total polyphenols

 The samples of freeze-dried potato were used for the extraction of polyphenols with 70%

aqueous acetone, as described by Nemś et al. (2015). Total polyphenol content was determined

using the Folin-Ciocalteu colorimetric method, as described by Gao, Bjork, Trajovski, & Uggla

(2000). Polyphenol content, expressed as milligrams gallic acid equivalent (GAE), was calculated

per 1 gram of DM.

2.4. Assay of amino acid composition

Freeze-dried samples, milled and sieved, were used for amino acid determination. The amino

acid composition was determined by ion-exchange chromatography after 23 hours’ hydrolysis with

6 N HCl at 110 °C. After cooling, filtering and washing, the hydrolyte was evaporated in a vacuum

evaporator at a temperature below 50 °C. The dry residue was dissolved in a buffer of pH 2.2. The

prepared sample was analysed using the ninhydrin method (Simpson, Neuberger, & Lin, 1976;

Spackman, Stein, & Moore, 1958). The pH 2.6, 3.0, 4.25, and 7.9 buffers were applied. The

ninhydrin solution was buffered at pH 5.5. The hydrolysed amino acids were determined using an

AAA-400 analyser (INGOS, Prague, Czech Republic). A photometric detector was used, working at

two wavelengths, 440 nm and 570 nm. A column of 350 × 3.7 mm, packed with ion exchanger

Ostion LG ANB (INGOS) was utilised. Column temperature was kept at 60–74°C and detector at

121 °C. The calculations were carried out relative to an external standard. No analysis of tryptophan

was carried out. During the 23 hours’ acid hydrolysis at 110 °C, Trp, Asn and Gln are totally lost

(Asn and Gln turn to Asp and Glu, respectively). The losses of Cys, Met, Thr, Ser and Tyr reach up

to 15%.

2.5. Expression of the results

The amino acid content in potatoes was calculated on a dry weight (DW) basis and the

composition of amino acids expressed on the nitrogen basis (g per 16 g N). Moreover, it was

necessary to compare the amino acid composition of the coloured-flesh potatoes to a reference

protein. The amino acid pattern for high-quality protein established by the Joint Food and

Agriculture Organisation/World Health Organisation (FAO/WHO) Committee in 1991, according

to Young and Pellett (1991), was chosen. Levels were calculated on the basis of the essential amino

acid composition of the chemical scores (CS), according to the Mitchell and Block method

(Osborne & Voogt, 1978) and the integrated EAA index (Oser, 1951).

2.6. Statistical analyses

All data were statistically analysed using Statistica 10.0 (Statsoft, Inc., Tulsa, OK).

Homogenous groups and least significant difference (LSD) values were denoted using Duncan’s

multiple comparison test. The significance level was set at α = 0.05, with one-way analysis of

variance (ANOVA) for three variables.

3. Results and discussion

3.1.Chemical and amino acid composition

The analysed coloured potato varieties were characterised by different DM, nitrogen and total

polyphenol contents (Table 1). The total and coagulable protein content in the potatoes depended on

the variety, not on the flesh colour. Purple-fleshed Blue Congo and red-fleshed Rote Emma

contained a greater total protein content than the potatoes of other varieties studied (2.87 g and 2.63

g 100 g‒1 fresh weight (FW), respectively). The lowest content of total protein was noted in tubers

of purple-fleshed Blaue Annelise and yellow-fleshed Fresco (2.01 g and 2.16 g 100 g‒1 FW,

respectively). There were comparatively smaller differences among the studied samples of potatoes

in respect of coagulable protein content. Potatoes of purple-fleshed Blaue Annelise and Blue

Congo, red-fleshed Rote Emma and yellow-fleshed Vineta varieties had similar coagulable protein

levels (from 0.55 g to 0.60 g 100 g‒1 FW). In contrast, the amount of coagulable protein was

significantly higher in red-fleshed Herbie 26 (0.70 g 100 g‒1 FW) and lowest in tubers of the

yellow-fleshed Fresco variety (0.34 g 100 g‒1 FW).

Jansen & Flamme (2006) studied 18 potato varieties/breeding clones of white- and purple-

fleshed cultivars and reported no distinct differences concerning DM, starch and protein content,

with comparable values to traditional white- or yellow-fleshed varieties. The total protein content is
said to depend mainly on the potato variety and fertilisation (Leszczyński, 2002; Mazur & Kreft,

1983; Rexen, 1976; Stankiewicz, Bombik, Rymsza, & Starczewski, 2008).

Purple-fleshed Blaue Annelise and Blue Congo varieties presented the highest content of total

polyphenols (2.50 mg and 1.84 mg g‒1 DM, respectively), whereas the lowest content was observed

in Fresco tubers (0.58 mg g‒1 DM). Potato varieties with coloured flesh contain more polyphenolic

compounds, including anthocyanins, which are not found in yellow- or cream-fleshed varieties.

According to Brown (2005), purple- and red-fleshed potato varieties contain at least twice the levels

of phenolic acid that yellow-fleshed potatoes contain.

Both the variety and the storage conditions affected the protein and amino acid content of the

analysed tubers of different flesh colours (Table 2). However, the biggest differences in the content

of these compounds were observed among samples stored for different times and among different

potato varieties, independently of flesh colour. There was no influence of storage temperature on

the coagulable protein contents of the evaluated tubers. The studies showed that as the storage time

increased, the total protein content (DM basis), the sum of all amino acid contents and that of all

EAA contents declined. The loss of nitrogen compounds at three months’ storage amounted to

about 6.4%, but at six months increased to 19% on average compared to the potato tubers before

storage. The coagulable protein content (DM basis) of the analysed potatoes increased at three

months storage by an average 25% that was maintained (3.61 g 100 g‒1 DW) at six months.

Decreasing the total protein content but increasing the coagulable protein content at three months’

storage can be explained by the synthesis of proteins occurring especially in the first months of

tuber storage (Brierley et al., 1996). On the other hand, increasing the total nitrogen and decreasing

the coagulable protein content at six months’ storage was a result of protein hydrolysis, as the

soluble nitrogen increased independently of temperature. According to Brierley et al. (1996),

protein degradation is associated with the end of tuber dormancy and the mobilization of nitrogen

reserves for sprout formation, while the breakdown of proteins is consistent with an increase in

proteolytic enzyme activity.

 The initial average of the sum of amino acids was 10.09 g 100 g ‒1 DW. At three months’

storage, it remained at 9.27 g 100 g‒1 DW, constituting 92% of the original amount. However, at six

months’ storage, this had decreased relative to the initial value by, on average, 28%, with a

maximum 7.29 g 100 g‒1 DW detected (Table 2). The total content of EAAs decreased, on average,

by 30% at six months relative to the samples before storage. Some previous authors (Brierley et al.,

1996; Stankiewicz et al., 2008) documented that a decrease in the content of amino acids in the DM

of tubers under the influence of storage time was strictly connected to the accompanying decline in

the total protein content. Additionally, an increase in total nitrogen concentration (DM basis) has

been correlated with a reduction in the EAA content of the total protein (Danilchenko, Pranaitiene,

Tarasieviciene, & Venskutoniene, 2008; Mitrus et al., 2003; Pęksa et al., 2013) and depends

primarily on the potato variety, but also on the fertilisation.

Moreover, from the results presented in Table 2, tubers of individual varieties differed

significantly in their content of nitrogen compounds, including total and coagulable protein, as well

as the sum of all and EAAs indices. Among all six potato varieties assessed, red-fleshed Herbie 26

and Rote Emma and yellow-fleshed Fresco tubers were characterised by higher total protein and

sum of all amino acids (DM basis) after six months’ storage, whereas purple-fleshed Blaue

Annelise variety had the lowest quantities of these compounds. Less diversity among samples of the

analysed tubers was evident regarding the coagulable protein content (DM basis), which ranged

from 3.04 g to 3.70 g 100 g‒1 DW. Galdón et al. (2010) observed relatively high variation in the

amino acid data for all the potato cultivars grown in the Canary Islands. Thus, the genetic

characteristics of the potato varieties decisively influenced the amino acid profile.

The data did not indicate a significant influence of storage temperature on the content of almost

all the determined amino acids. Potatoes stored at 2 ºC contained (DW basis) slightly more amino

acids, mainly asparagine, glutamine, proline, leucine, lysine and arginine, than tubers stored at 5 ºC.

However, there was a statistically significant difference for the amino acids valine and isoleucine.

The tuber storage temperature did not impact on the serine, glycine, cysteine, tyrosine and
phenylalanine content. Likewise, Talley, Toma, and Orr (1984) stated that observed differences in

the content of particular amino acids in potatoes stored at different temperatures (3.3 °C and 7.2 °C)

were not significant. Moreover, these authors did not find significant differences for methionine,

isoleucine and tyrosine content. However, content of asparagine, threonine, serine, proline, glycine,

valine, leucine, phenylalanine, histidine, lysine, arginine and tryptophan increased during storage at

3.3 °C, while glutamine decreased.

Throughout the six months’ storage, losses were observed in the majority of the potato amino

acids (DM basis) (Table 2), both EAAs and non-essential amino acids (NEAAs). The storage time

influenced the content of the individual amino acids more than the storage temperature, the

differences at six months’ storage amounting to 21–38% relative to non-stored tubers. Furthermore,

an increase (33%) in proline content occurred at three months’ storage. The storage time most

affected the content of threonine, valine, arginine, leucine, histidine, lysine, isoleucine and

asparagine. At three months’ storage, the examined tubers contained 6–18% less threonine, valine,

methionine, isoleucine, leucine, tyrosine and lysine (EAAs) and, among the NEAAs, there were

smaller amounts of asparagine, serine, glutamine, histidine and arginine. Losses of the remaining

amino acids in the first period of storage were smaller than 16%. At six months’ storage of tubers of

coloured flesh varieties, the content of individual amino acids (DM basis) decreased from 38% to

21% compared to the samples before storage. The smaller losses were observed for serine,

glutamine, asparagine NEAAs and tyrosine or isoleucine EAAs; higher declines were noted for

lysine, valine, leucine, threonine, histidine and arginine content. The amino acids content (DM

basis) of tubers during long-term storage decreased along with the total nitrogen content. This

relationship concurs with Eppendorfer and Eggum (1994), who assessed the quality of potatoes with

traditional light-coloured flesh. The authors stated a close association between nitrogen content and

the concentration (DM basis) of methionine, cysteine, threonine and lysine. According to Talley et

al. (1984), the concentration of individual amino acids typically followed the same order as the

nitrogen content. However, in many instances where the order was not followed, the values were
not significantly different. Stankiewicz et al. (2008) found that at seven months’ storage of Irga and

Ekra potato tubers, the total protein content significantly decreased.

There was significant differentiation among the contents of particular amino acids depending on

the potato variety (Table 2). Yellow-fleshed Fresco potatoes contained more EAAs, such as

threonine, methionine, tyrosine and phenylalanine, while among the NEAAs, asparagine, serine and

proline contents were notably higher than in the other analysed tubers. The lowest values of amino

acids, such as threonine, valine, lysine, asparagine and arginine, were found in purple-fleshed Blaue

Annelise potatoes. Significant amounts of most amino acids were detected in potatoes of Rote

Emma, Herbie 26 and Blue Congo varieties.

The content of total protein (DM basis) of analysed tubers of Blaue Annelise, Blue Congo, Rote

Emma, Vineta and Fresco varieties decreased during the six months’ storage, whereas in Herbie 26

no significant changes in the total nitrogen compounds occurred (Table 3). In potatoes of Blue

Congo, Rote Emma and Vineta varieties, differing in flesh colour, decreasing total protein content

was found at both three months’ and six months’ storage. Conversely, a significant decrease in total

protein content was noticed only at three months’ storage in Fresco variety and six months’ storage

in Blaue Annelise tubers. The highest loss of total nitrogen compounds at six months’ storage was

found in the yellow-fleshed Vineta variety, which reached 38%; the smallest losses (average 1–8%)

were recorded in red-fleshed Herbie 26 and yellow-fleshed Fresco varieties.

The content of coagulable protein (DM basis) of the potato tubers significantly increased at

three months’ storage, and changed slightly or plateaued during the next three months of storage

(Table 3). The proportion of coagulable protein (DM basis) increased maximally in the Fresco

variety (by about 42%) and minimally in Herbie 26 tubers (by about 6%).

The presented studies showed that storage for six months contributed to a gradual decrease in

the total amino acids (DM basis) of almost all potato samples, regardless of flesh colour (Table 2),

and ranged from 22% to 40%. Only red-fleshed Herbie 26 variety, characterised by an insignificant

increase in the total amino acids content at three months’ storage and at six months, retained the
same value as that of the respective tubers before storage. Černá and Kráčmar (2010) determined

that the total quantity of amino acids in potatoes decreased by around 9% to 28% at 16 weeks’

storage, depending on the potato variety.

There was observed an increase in dry matter content of most analysed potatoes. However, dry

matter of Blue Congo and Herbie 26 did not change significantly during storage. The increase of

dry matter content probably was the result of excessive evaporation of water from tubers stored in a

proportionally low humidity (85 ± 2%).

3.2.Nutritive quality of the protein

Potato protein is characterised by a high biological value, which is a measure of the proportion

of protein from a specific food that can be utilised to synthesise the proteins of the organism. The

nutritional value of proteins can be expressed by determining various indicators; for example, the

essential amino acids index (EAAI) and the CS. The CS is a comparison of each EAA in a specific

protein to the content of a standard protein, typically whole egg. The limiting EAA in the test

protein is expressed as the percentage of the amount of the same amino acid in the reference

protein. The CS of the EAAs and the EAAI were calculated with respect to the reference protein of

the joint FAO/WHO (1991), taking into account all EAAs besides tryptophan, the storage

temperature and time, as well as potato tuber variety.

The storage temperature of the six potato tubers of coloured flesh varieties did not significantly

influence the nutritional value of the protein, expressed as CS and EAAI (Table 4). In samples

stored at 2 °C, the average CS was 74 and was identical for sulfur amino acids and leucine. The

second limiting amino acid was threonine (CS = 85). In potatoes stored at 5 °C, the CS was 69–70

and concerned leucine and sulfur amino acids.

The storage time significantly affected the nutritive value of the studied samples of differing

flesh colour varieties (Table 4). The studies showed that the nutritional value indices of the protein

contained in potatoes, both CS and EAAI, decreased at three and six months’ storage. Only the CS

values of amino acids valine, phenylalanine and tyrosine, and isoleucine changed, but only slightly,
maintaining high levels. Similarly, Galdón et al. (2010) established the CS of all amino acids with

respect to the reference protein of the joint FAO/WHO (1991) calculated for each cultivar and

stated that the highest CS values were determined for aromatic amino acids (phenylalanine with

tyrosine) and for branched amino acids, like isoleucine, leucine and valine.

Before storage, the protein of the assessed potato samples characterised by CS, ranged from 84

to 156, whereas EAAI averaged 112. The limiting amino acids, before beginning the experiment,

were leucine (CS = 84) and methionine + cysteine (CS = 92). At three months’ storage, the CS

values ranged from 67 (for sulfur amino acids) to 147 (for aromatic amino acids). Extending the

storage time to six months contributed to further lowering the nutritional value of the protein

contained in the analysed tubers of the six potato varieties. This behaviour was reflected in the

EAAI, which decreased by over 30% relative to the EAAI of the potatoes before storage. The CS

(55 and 65 for sulfur amino acids and leucine, respectively) also limited the potato quality.

The analysed potato tubers of the different coloured flesh varieties showed significant

differentiation in terms of the nutritional value of the protein contained regardless of storage

conditions (Table 3). On average, Fresco, Rote Emma and Herbie 26 varieties were characterised by

the highest EAAI of protein, with values of 118, 110 and 96, respectively; leucine (CS = 84),

methionine + cysteine (CS = 78), and leucine (CS = 72) were the amino acids limiting the protein

quality of these potato varieties.

The nutritional value of the potato varieties, expressed as CS relative to the referenced standard

protein (FAO/WHO, 1991), changed depending on the storage time and variety (Figure 1). Before

storage, the yellow-fleshed Fresco potatoes contained full-valued protein (CS = 142), which

contrasted with the tubers of the other varieties, wherein the amino acid limiting the protein quality

was leucine, and CS, which ranged between 67 and 85. Our results corroborate the data published

by Eppendorfer and Eggum (1994), who found that methionine + cysteine and leucine were the

most limiting amino acids in the studied potato tubers. Conversely, Sotelo, Contreras, Soussa, and
Hernández (1998) reported that the sulfur amino acids were the limiting amino acids in all

investigated potato varieties.

The nutritional value of protein contained in potatoes of the evaluated varieties decreased

during storage, depending on the variety. This was particularly the case for purple-fleshed Blue

Congo and yellow-fleshed Fresco potatoes at six months’ storage, and for Blaue Annelise at three

months’ storage (Figure 1). At three months’ storage, the CS of the protein nutritional value

decreased in these varieties by, on average, over 30%; in tubers of the Blue Congo and Fresco

varieties, it decreased by a further 12–18% at six months. Leucine was the amino acid limiting the

quality of protein of stored potatoes of the Fresco variety (CS range 69–57). Conversely, Blue

Congo was limited by methionine + cysteine (CS range 49–43), while the quality of Blaue Annelise

protein was limited by methionine + cysteine at three months’ storage (CS = 54) and threonine at

six months (CS = 70). Relatively small changes in the nutritional value at six months’ storage in

comparison to their non-stored counterparts were observed for the protein of red-fleshed Herbie 26

and yellow-fleshed Vineta varieties. The nutritional value of protein of these potatoes, expressed as

CS, maintained a similar level to the samples before storage. At three months’ storage, the limiting

amino acids were, proportionally, leucine (CS = 58) and methionine + cysteine (CS = 59); at six

months, these were methionine + cysteine (CS = 65) and leucine (CS = 71).

At six months’ storage, the CS of all analysed varieties was 43–71. However, the protein of

Vineta and Blaue Annelise tubers presented the highest nutritional value, despite containing less

total nitrogen compounds than the other tubers. The data are consistent with van Gelder and Vonk

(1980), who studied the amino acid composition of the protein of 34 potato varieties and stated that

the slight variation in the amino composition of coagulable protein hardly affects the EAAI.

Additionally, the amino acid composition of coagulable potato protein was largely the same in

varieties with low and high protein content. Similar findings were also confirmed by our previous

research (Pęksa et al., 2013) and by other authors (Eppendorfer & Eggum, 1994; Rexen, 1976).
4. Conclusion

The presented study found that during storage of the six potato samples of varieties with

different coloured flesh (DM basis), the nitrogen compounds decreased, along with the amino acid

content (except proline). However, an increase in the coagulable protein content occurred, to the

greatest extent in tubers of the yellow-fleshed Fresco variety (by about 42%) and to the least extent

in the DM of red-fleshed Herbie 26 tubers (by about 6%). The storage temperature did not

significantly influence the coagulable protein content and only insignificantly affected some of the

amino acid content: samples stored at a lower temperature (2 ºC versus 5 ºC) contained less valine

and isoleucine, for instance. The storage time mostly impacted the content of amino acids, such as

threonine, valine, arginine, leucine, histidine, lysine, isoleucine and asparagine. Among the samples

of tubers of the six potato varieties, there was lower diversity regarding the coagulable protein (DM

basis) than the total protein and the sum of all amino acid content. The highest losses of total

nitrogen compounds, following six months’ storage, were found in yellow-fleshed potatoes of the

Vineta variety, which reached 38%; in tubers of red-fleshed Herbie 26 and yellow-fleshed Fresco

varieties, however, this value was lower than 8%.

The increase in the coagulable protein content (DM basis) of potatoes during long-term storage

did not impact its increased quality, as confirmed by the decreasing values of the nutritional value

indicators (EAAI, CS) determined at three and six months of storage, independently of potato

variety and flesh colour. Before storage, the limiting amino acid was leucine (CS = 84), but at three

and six months’ storage, it was the sulfur amino acids, with CS values of 67 and 55, respectively.

After storage at 2 ºC and 5 ºC, leucine and sulfur amino acids limited the protein quality of the

studied potatoes. In samples stored at 2 ºC, the average CS was 74; in potatoes stored at 5 ºC, it was

69–70. After six months of storage, the CS of all analysed varieties ranged between 43 and 71.

However, the proteins of Vineta and Blaue Annelise were characterised by the highest nutritional

value, despite containing fewer total nitrogen compounds than the other tubers.
Acknowledgements

This project was financed by the National Science Centre, granted on the basis of decision

DEC-2013/11/N/NZ9/00117.

This publication was supported by the Wroclaw Centre for Biotechnology programme at the

Leading National Research Centre (KNOW) for the years 2014–2018.

The authors declare no commercial or financial conflict of interest.

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 0-month
120
100 F
BA RE
80
H26 BC
60
V
40
20
0
0 2 4 6 8 10 12 14
Total protein (g·100g-1 DW)

3-month
120
100
RE
80 F
V H26
60 BA
40
20 BC
0
0 2 4 6 8 10 12 14
Total protein (g·100g-1 DW)

6-month
120
100
80 V H26
60 BA
BC F
40 RE
20
0
0 2 4 6 8 10 12 14
Total protein (g·100g-1 DW)

Fig 1. CS values of the protein contained in potatoes of different flesh colour varieties after 0, 3 and 6-month
storage in relation to total protein content.
The first limited amino acid: (0-month) Leu; (3-month) Met+Cys (BA, BC, RE, V), Leu (H26, F); (6-month) Thr (BA),
Met+Cys (BC, H26, RE), Leu (V, F)
Table 1. Chemical composition of experimental potatoes at the beginning of storage

variety colour of dry matter total protein coagulable total

flesh protein polyphenols
g·100g‒1 FW g·100g‒1 FW g·100g‒1 FW mg·g‒1 DM
d f b a
Blaue Annelise purple 20.55 ±0.22 2.01 ±0.01 0.61 ±0.03 2.50 ±0.08
a a b b
Blue Congo 23.57 ±0.71 2.87 ±0.01 0.61 ±0.02 1.84 ±0.08

Herbie 26 red 22.05b±0.05 2.45c±0.02 0.70a±0.05 1.21c±0.20

Rote Emma 21.42c±0.35 2.63b±0.06 0.60b±0.01 0.83d±0.09

Vineta yellow 19.92d±0.15 2.35d±0.01 0.55b±0.02 0.66e±0.07

e e c f
Fresco 17.34 ±0.24 2.16 ±0.01 0.34 ±0.01 0.58 ±0.08

Values are mean ± SD of three determinations;

a,b,c,d,e,f – means in a column with the same letter are not significantly different (p < 0.05)
Table 2
Amino acid concentration and protein content (g·100 g‒1 DW) in potatoes of different flesh colour varieties
as influenced by storage conditions and variety.

storage storage time variety

temperature (month)
(°C)
2 5 0 3 6 BA BC H26 RE V F
a b a b c d c b c c a
2.15 ±0. 2.08 ±0. 2.43 ±0. 2.18 ±0. 1.73 ±0. 1.77 ±0. 2.07 ±0. 2.32 ±0. 2.08 ±0. 2.06 ±0. 2.39 ±0.
Asp 39 49 19 41 36 37 57 22 33 51 35
b a a b c d c c b c a
Thr 0.31 ±0. 0.32 ±0. 0.39 ±0. 0.32 ±0. 0.24 ±0. 0.26 ±0. 0.28 ±0. 0.30 ±0. 0.34 ±0. 0.30 ±0. 0.42 ±0.
* 09 10 10 05 05 07 08 03 06 08 01
a b c c c b b c a
0.34± 0.34± 0.38 ±0. 0.35 ±0. 0.30 ±0 0.31 0.30 ±0. 0.38 ±0. 0.36 ±0. 0.31 ±0. 0.41 ±0.
Ser 4
0.07 0.07 04 08 .06 ±0.0 08 06 06 07 06
a b a b c d b c a d c
1.50 ±0. 1.34 ±0. 1.58 ±0. 1.48 ±0. 1.20 ±0. 1.16 ±0. 1.60 ±0. 1.46 ±0. 1.66 ±0. 1.20 ±0. 1.44 ±0.
Glu
32 29 26 29 27 17 41 18 29 21 24
a b b a b bc d b bc c a
0.40 ±0. 0.33 ±0. 0.31 ±0. 0.46 ±0. 0.32 ±0. 0.35 ±0 0.30 ± 0.37 ±0. 0.35 ±0 0.33 ±0. 0.47 ±0.
Pro
16 08 07 17 06 .13 0.06 08 .02 14 21
a a b d d c a cd b
0.3± 0.3± 0.34 ±0. 0.33 ±0. 0.24 ±0. 0.26 ±0. 0.26 ±0. 0.30 ±0. 0.39 ±0. 0.27 ±0 0.34 ±0.
Gly
0.09 0.09 07 11 04 04 07 04 13 .05 10
a b a a b c c c a d b
0.28 ±0. 0.27 ±0. 0.29 ±0. 0.29 ±0. 0.25 ±0. 0.26 ±0. 0.25 ±0. 0.26 ±0. 0.34 ±0. 0.23 ±0. 0.30 ±0.
Ala
07 05 04 08 05 04 05 03 08 02 06
a c b b b a a b a
Cys 0.05± 0.05±0.0 0.06 ±0. 0.04 ±0. 0.05 ±0. 0.05 ±0. 0.05 ±0. 0.06 ±0. 0.06 ±0. 0.05 ±0. 0.06 ±0.
* 0.01 1 01 02 01 02 02 01 02 01 02
ami Val
b
0.45 ±0.
a
0.48 ±0.
a
0.53 ±0.
b
0.50 ±0.
c
0.35 ±0f
e
0.35 ±0.
c
0.43 ±0.
b
0.47 ±0.
a
0.55 ±0.
d
0.39 ±0.
a
0.58 ±0.
no * 17 13 14 16 .07 06 12 10 17 09 18
acid a b a b c c c b b c a
Me 0.15 ±0. 0.14 ±0. 0.19 ±0. 0.14 ±0. 0.10 ±0. 0.12 ±0. 0.11 ±0. 0.15 ±0. 0.15 ±0. 0.12 ±0. 0.19 ±0.
t* 06 05 05 04 02 05 04 03 04 04 07
b a a b c d c b a d a
0.32 ±0. 0.34 ±0. 0.38 ±0. 0.33 ±0. 0.27 ±0. 0.28 ±0. 0.31 ±0. 0.34 ±0. 0.39 ±0. 0.27 ±0. 0.37 ±0.
Ile*
10 08 08 10 04 05 08 05 10 05 13
a b a b c d d c a d b
Leu 0.53 ±0. 0.49 ±0. 0.60 ±0. 0.53 ±0. 0.40 ±0. 0.44 ±0. 0.44 ±0. 0.51 ±0. 0.63 ±0. 0.45 ±0. 0.60 ±0.
* 17 13 12 16 07 09 12 07 18 08 19
a b c d c c b d a
Tyr 0.32± 0.31±0.0 0.36 ±0. 0.31 ±0. 0.27 ±0. 0.27 ±0. 0.31 ±0. 0.31 ±0. 0.35 ±0. 0.28 ±0. 0.37 ±0.
* 0.09 6 07 08 04 05 08 05 08 06 09
a a b d c c b d a
Phe 0.66±0.2 0.64±0.1 0.70 ± 0.69 ± 0.55 ± 0.53 ±0 0.60 ±0. 0.60 ±0. 0.77 ±0. 0.50 ±0. 0.87 ±0.
* 2 8 0.23 0.22 0.11 .08 12 10 23 08 24
a± b a b c d c b a d b
0.20 0. 0.19 ±0. 0.22 0.20 ±0. 0.15 ±0 0.16 ±0. 0.18 ±0. 0.21 ±0. 0.23 ±0. 0.16 ±0. 0.21 ±0.
His
06 04 ±0.03 06 .03 03 04 04 06 03 06
a b a b c d c b a c a
Lys 0.55 ±0. 0.51 ±0. 0.62 ±0. 0.55 ±0. 0.43 ±0. 0.41 ±0. 0.48 ±0. 0.55 ±0. 0.63 ±0. 0.47 ±0. 0.64 ±0.
* 18 13 14 17 08 07 10 09 19 09 19
a b a b c e b b a d c
0.58 ±0. 0.55 ±0. 0.68 ±0. 0.57 ±0. 0.45 ±0. 0.36 ±0. 0.66 ±0. 0.64 ±0. 0.82 ±0. 0.41 ±0. 0.51 ±0.
Arg
22 20 20 21 16 06 24 11 15 12 12
a b a b c e c b a d a
9,15 ±1, 8.61 ±1. 10.09 ±1 9.27 ±1. 7.29 ±1. 7.33 ±1. 8.64 ±2. 9.22 ±1. 10.10 ±1 7.81 ±1. 10.19 ±1
∑aa 98 88 .45 84 32 09 09 09 .85 57 .99
b a b c e d c b de a
3,39±1,0 3.22 ±0. 3.85 ±0. 3.41 ±0. 2.66 ±0. 2.70 ±0. 3.02 ±0. 3.29 ±0. 3.88 ±1. 2.84 ±0 4.10 ±1.
∑eaa 4 83 91 96 45 45 74 52 02 .56 24
a b a b c e c b b d a
10,81 ±1 10.44 ±1 11.6 ±0. 10.86 ±1 9.42 ±1. 8.81 ±1. 10.24 ±1 11.47 ±0 11.53 ±0 9.94 ±2. 11.77 ±0
TP ,63 .81 95 .45 89 69 .52 .81 .74 12 .91
b a c cd a b c d
3,33±0,6 3.28±0.6 2.70 ±0. 3.61 ±0. 3.61a±0 3.24 ±0. 3.13 ±0 3.70 ±0. 3.58 ±0. 3.16 ±0. 3.04 ±0.
CP 6 9 38 62 .56 38 .41 43 76 88 85
Values are means ±SD of six determinations.. a,b,c,d,e – means in a row with the same letter are not significantly different (p < 0.05)
* Essential amino acid; ∑aa – sum of amino acids; ∑eaa - sum of essential amino acids; TP - total protein; CP - coagulable protein;
potato varieties: Blaue Annelise (BA), Blue Congo (BC), Herbie 26 (H26), Rote Emma (RE), Vineta (V), Fresco (F)
Table 3

The content of protein, the sum of amino acids (g·100 g‒1 DW) in potatoes of six varieties and their dry
matter (g·100 g‒1 FW) as influenced by storage time and variety (mean values of the storage temperature).

attrib vari Blaue Annelise Blue Congo Herbie 26 Rote Emma Vineta Fresco
ute ety

time
of
stor 0 3 6 0 3 6 0 3 6 0 3 6 0 3 6 0 3 6
age

(mo
nth)

total 9.7 9.5 7.11 12. 9.70 8.8 11.1 12. 11.0 12. 11.5 10. 11. 10. 7.27 12. 11. 11.
h h j b h i e ef d g g j j d d
protei 7 6 19 3 2 27 3 26 1 83 78 77 48 36 46
ab ab c
n ±0. ±0.1 ±0. ±0.4 ±0. ±0.1 ±0.7 ±0.4 ±1. ±0.1 ±0. ±1. ±0.
07 ±0. 7 04 3 40 2 ±0. 2 ±0. 6 ±0. 25 2 10 08 91
90 80 29 ±0. 02
61

coagu 2.8 3.2 3.58 2.6 3.39 3.3 3.17 3.3 4.00 2.8 3.84 4.1 2.8 3.8 2.84 1.9 3.4 3.7
h fg de i ef ef g ef ab hi bc a h bc h j ef cd
lable 4 9 0 9 9 0 2 5 0 9 3 2
protei ±0. ±0.3 ±0.2 ±0. ±0. ±0.2 ±0. ±0.3 ±1. ±0.2 ±0. ±0.
16 ±0. 2 ±0. 7 10 ±0.2 15 5 10 4 ±0. 10 0 ±0. 46 39
n 20 10 2 50 ±0. 09
10

Σaa 8.6 7.0 6.30 11. 7.98 6.8 8.49 10. 8.74 10. 11.4 8.2 9.7 8.3 5.78 12. 10. 7.8
e h i b fg h e c e c b d ef j
8 2 13 0 45 60 5 7 2 7 34 38 3g
efg a c
±0. ±0. ±0.5 ±0. ±0.4 ±0. ±0. ±0. ±0.9 ±0. ±0.
07 30 2 46 4 20 ±0.3 79 ±0.7 09 0 ±0. 39 ±0.1 ±0. ±0. 21
6 8 ±0. 23 7 76 72
99

Σeaa 3.2 2.5 2.34 3.9 2.68 2.4 3.04 3.8 2.95 3.7 4.82 3.0 3.4 2.9 2.17 5.6 3.6 3.0
ef hi hi c gh hi fg c fg cd b fg de fg i a cd fg
4 3 5 2 9 8 3 3 2 4 4 4
±0.4 ±0. ±0.1 ±0.2 ±0. ±0. ±0. ±0.1 ±0. ±0.
±0. ±0. ±0.3 4 11 4 ±0. 0 09 ±0.8 ±0. 07 22 1 ±0. 41 12
13 10 5 ±0. 45 1 49 82
16

dry 20. 23. 24.1 23. 23.1 24. 22.0 24. 23.7 21. 22.0 25. 19. 20. 21.9 17. 20. 24.
ef ab bcd bc cde ab cde ab abc de cde a f ef cde g f ab
matte 55 06 8 57 7 68 5 21 1 42 2 51 92 85 1 34 03 06
f
r ±0. ±0. ±0.7 ±0. ±0.9 ±1. ±0.0 ±0. ±0.9 ±0.9 ±1. ±0. ±0. ±1.1 ±0. ±0. ±1.
22 66 8 71 9 12 5 81 8 ±0. 8 13 15 54 8 24 89 31
35

Values are mean ±SD of three determinations;

a,b,c,d,e,f, g,h,i,j – means in a row with the same letter are not significantly different (p < 0.05)
 Table 4
Essential amino acid index (EAAI) and chemical scores of essential amino acids present in potatoes of
different flesh colour varieties in relation to temperature, time of storage and variety.
essential amino acids
factor EAAI Thr Met+Cys Val Ile Leu Phe+Tyr Lys
storage 2 96 85 74 119 106 74 144 88
temperature
(°C) 5 95 87 70 127 112 69 140 81

storage 0 112 106 92 140 125 84 156 99

time
3 97 87 67 132 109 74 147 88
(month)
6 76 65 55 93 89 56 121 69

variety Blaue Annelise 79 71 63 93 92 63 118 65

Blue Congo 86 76 59 114 102 63 134 77

Herbie 26 96 82 78 125 112 72 134 88

Rote Emma 110 93 78 146 129 88 165 101

Vineta 82 82 63 103 89 63 115 75

Fresco 118 115 92 154 122 84 182 102

protein standard FAO/WHO (1991) 3.4 2.5 3.5 2.8 6.6 6.3 5.8
(g/16 g N)
HIGHLIGHTS

>Amino acids profile and protein quality of stored coloured potatoes were analysed

>Storage temperature did not affected protein content and its nutritive quality

>Losses of Lys, Val, Leu, Thr, His and Arg amino acids occurred in stored potatoes

>The protein of stored potatoes of Fresco and Herbie 26 varieties was of the highest quality

28